Environmental Management

DOI 10.1007/s00267-015-0562-5

Livestock Predation by Puma (Puma concolor) in the Highlands

of a Southeastern Brazilian Atlantic Forest
Francesca Belem Lopes Palmeira1,3 • Cristiano Trapé Trinca1,3 • Claudio Maluf Haddad2

Received: 4 February 2014 / Accepted: 8 June 2015

Ó Springer Science+Business Media New York 2015

Abstract We evaluated local opinion about reducing
livestock losses to puma (Puma concolor) and the potential
for conflict among livestock breeders inside a protected
area in the highlands of a southeastern Brazilian Atlantic
forest. We also quantified the number and type of livestock
losses, and determined if predation by puma was correlated
with property profile and landscape characteristics. We
conducted semistructured interviews with 42 livestock
breeders sampled in 36 rural properties. When asked how
to reduce predation, 33 % of livestock breeders refused to
answer, 26 % suggested improving livestock husbandry
practices, 19 % stated that there was no appropriate action,
17 % favored removing the ‘‘problem’’ individual, and 5 %
suggested killing the puma. Opinion on how to solve pre-
dation was independent of herd size and history of losses,
and was correlated with respondent age class. Older
respondents tended to suggest removing or killing pumas.
Attitudes toward predation represented high potential for
conflict among livestock breeders who demonstrated high
discordance among responses. Horses were the most
common prey (51 %), followed by cattle (28 %), sheep
(17 %), and goats (4 %); totaling 47 animals attacked
between 2004 and 2007. Annual predation was
& Francesca Belem Lopes Palmeira
yagouaroundi@yahoo.com.br;
http://www.jaguarjuruena.com
1
Research Department, Reserva Brasil, Av Dr Silva Melo 520,
Apto 606, Jd Taquaral, São Paulo, SP CEP 04675-010, Brazil
2
Department of Animal Science, ‘‘Luiz de Queiroz’’ College
of Agriculture, University of Sao Paulo (ESALQ/USP), Av
Pádua Dias 11, Piracicaba, SP CEP 13418-900, Brazil
3 lynx), jaguars (Panthera onca), pumas (Puma concolor),
Present Address: Jaguar Juruena, R Joao Braz Alves, QD 04,
Lote 16, Japuranã, Nova Bandeirantes, MT CEP 78565-000,
Brazil
approximately 12 ± 5 animals, equivalent to 0.4 % of the
total livestock. Property elevation and distance from the
urban center were the main predictors of predation proba-
bility. This survey used a novel approach that has not been
addressed directly in other studies on livestock predation
and demonstrated that the high potential for conflict among
livestock breeders should be considered before imple-
menting management actions.
Keywords Attitudes  Conservation  Generalized linear
model  Human–wildlife interaction  Potential for conflict
index
Introduction
The loss of livestock due to apex predators is a worldwide
issue and involves both ecological and social factors of
human–wildlife conflicts (Graham et al. 2005). Although
‘‘human–wildlife conflict’’ is a widely accepted term to
describe negative interactions between wild animals and
people, perhaps a more appropriate term for the losses caused
by wildlife is human–wildlife relationship or human–wild-
life interaction. Indeed, aspects of human–wildlife conflict
involve disagreements over how wildlife should be managed
(i.e., human–human conflict) (Peterson et al. 2010).
Big cats are responsible for attacks on livestock
throughout their distribution, and the severity of conflict
increases with felid body mass, which means higher levels
of conflict for larger species such as caracals (Caracal
caracal), cheetahs (Acinonyx jubatus), Eurasian lynx (Lynx
snow leopards (Uncia uncia), leopards (Panthera pardus),
lions (Panthera leo), and tigers (Panthera tigris) (Inskip

123

and Zimmermann 2009). However, although there is
enormous potential for big cats to attack livestock, mor-
tality by other causes (e.g., diseases, malnutrition, mal-
practice, and others) has a higher impact on livestock
losses than predation (Oli et al. 1994; Mazzolli et al. 2002;
Palmeira et al. 2008).
Among the Brazilian biomes inhabited by jaguar and
puma, the Atlantic Forest has suffered most from fragmen-
tation processes, with only 11–16 % of the original forested
area remaining (Ribeiro et al. 2009). This remaining area still
maintains high levels of biodiversity and endemism (Mit-
termeier et al. 1999). Although 88 % of the original forest is
completely depleted of four species of large mammals
[jaguar, tapir (Tapirus terrestris), white-lipped peccary
(Tayassu pecari), and muriqui (Brachyteles arachnoides)],
55 % is considered unsuitable for them (Jorge et al. 2013).
Even pumas, which appear to be more tolerant of forest loss
and fragmentation than jaguars (De Angelo et al. 2011), have
been affected by loss of genetic variation (Miotto et al. 2011;
Castilho et al. 2012). There is evidence that hunting, and not
lack of landscape connectivity, is a major threat for this
species (Castilho et al. 2011). Although illegal, the killing of
pumas as a result of livestock predation is carried out in
ranches across the biome (Guix 1997; Mazzolli et al. 2002;
Verdade and Campos 2004). Consequently, pumas exhibit
differences in activity patterns among areas with different
hunting pressures (Paviolo et al. 2009). When there is little
hunting and high tolerance by humans, puma populations
achieve high densities even in a modified landscape (Maz-
zolli 2010). Pumas consume a diet based on a broad range of
prey and exhibit high behavioral plasticity in using anthro-
pogenic resources from the agricultural matrix (Magioli et al.
2014), which should predispose the species to predating on
livestock.
Public attitudes toward livestock losses due to big cats
have become less tolerant, to the point where there are very
few surviving populations outside of protected areas
worldwide (Jackson and Nowell 1996), thus increasing the
conflict with wildlife conservation goals. To reduce hunt-
ing as retaliation for livestock predation, several strategies
have been discussed on a regional scale including predator
control (Linnell et al. 1997), improving livestock hus-
bandry practices (Ogada et al. 2003; Kissling et al. 2009),
paying financial compensation (Bulte and Rondeau 2005;
Maclennan et al. 2009; Dickman et al. 2011), and devel-
oping environmental education (Holmern et al. 2007).
While there is a generally good understanding of the
dynamics of human–wildlife conflicts, this understanding
can vary among sites depending on the local determinants.
This could explain why there is no consistent or reliable
framework for managing predation (Graham et al. 2005). It
should be necessary to combine baseline research, partic-
ipatory planning, and monitoring to engage affected
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Environmental Management
communities and wildlife managers in the co-management
of human–wildlife conflicts (Treves et al. 2006). Attitudes
toward predation may be affected by a wide range of
personal and sociocultural aspects (Marchini and Mac-
donald 2012; Soto-Shoender and Main 2013) leading to
high disparity in opinions. Indeed, this conflicting opinion
among stakeholders is a critical—yet hitherto largely
unrecognized—first step to address for any successful
predator management and mitigation program.
Based on these observations, our goals were (1) to eval-
uate the determinants of local opinion in relation to personal
profile, herd size, and history of livestock losses to puma and
the potential for conflict among livestock breeders, (2) to
quantify the number and type of livestock losses to pumas,
and (3) to determine if predation was correlated with prop-
erty profiles and landscape characteristics in properties sur-
rounding a protected area in the highlands of a southeastern
Atlantic forest. The survey represents a novel approach that
has not been addressed directly in other studies on livestock
predation. This is highly relevant as there is a current push to
analyze human–wildlife conflicts from a human dimensions
and psychological perspective. Finding good predictors of
the likelihood of livestock losses to pumas and attitudes
toward predation can help us develop preventive strategies to
mitigate the potential for conflict.
Study Area
This study was carried out surrounding the Campos do Jor-
dão State Park (CJSP), a protected area which is located
between 22°370 –22°450 S and 45°240 –45°310 W, bordered by
São Paulo and Minas Gerais states, southeastern Brazil
(Fig. 1). It is part of the Serra da Mantiqueira Environmental
Protection Area (APA) composed of some parks and several
private properties, totaling an area of 421,804.46 ha. The
vegetation consists of high-altitude grasslands and Araucaria
forest (Araucaria angustifolia), which are typical ecosys-
tems of high and cold regions of the Atlantic forest biome.
CJSP covers an area of 8341 ha and the elevation ranges
from 1030 to 2007 m. The climate is subtropical (upland),
with a dry winter and mild summer (cwb) as classified by
Köeppen. The average temperature varies from 27.2 °C in
the summer to -4.4 °C during the winter. Rainfall varies
from 1500 to 2000 mm and occurs mostly during the spring
and summer (Seibert et al. 1975; Clauset and Soares 1999).
CJSP was one of the first regions to be settled in
southeastern Brazil. In 1975, the park’s management plan
already reported changes to the vegetation resulting from
anthropogenic activities in the region (e.g., selective log-
ging, reforestation with exotic conifers, extensive pastures,
management of fields with fire and disorderly occupation;
Seibert et al. 1975).
Environmental Management

Fig. 1 Location of respondents (n = 42) and reported predation on livestock in the Campos do Jordão State Park (CJSP) and Serra da
Mantiqueira Environmental Protection Area (APA), southeastern Brazil

Methods livestock breeders (from May to August 2007) in an

Data Collection
From July 2006 to August 2007, we undertook nine field
expeditions, totaling 80 days of sampling effort. We con-
ducted interviews 10 months after first contact with
attempt to establish trust between the interviewer and the
interviewee, and provide more consistent data collection
(Oli et al. 1994; Palmeira and Barrella 2007).
We selected all breeders who reared livestock in the
park and surrounding properties, totaling 42 respondents
sampled in 36 properties. We only interviewed one

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respondent per property (n = 35), with the exception of the
park (n = 1) where there were three employees responsible
for livestock kept in the area. We also interviewed four
herders who had no property, but kept their livestock loose
in the park or on leased pastures of the surrounding prop-
erties. We conducted one interview per interviewee, lasting
an average of 26 ± 13 min, with a minimum of 9 and
maximum of 68 min. The interviews were held face to face
at the main house of each property and conducted in Por-
tuguese, which is the native language of interviewees and
interviewers.
We divided the questionnaire into three sections: (1)
profiles of properties and respondents; (2) livestock pre-
dation; and (3) local opinion (Table 1). During sections (1)
and (2), we used structured questions with precise wording
and fixed response categories. To avoid influencing the
replies in section (3), we asked semistructured questions,
which enabled respondents to express their opinions with
spontaneous responses. To obtain the property profile, we
recorded each property’s purpose and size (ha) and used
GPS to extract the geographic location (X and Y) of the
sites where livestock were kept. To obtain the respondent
profiles, we recorded each respondent’s age, the length of
residence in the region, place of birth, and their main
economic activity. To assess the damage caused by pre-
dation, we asked questions regarding the number and type
of livestock reared by respondents, husbandry practices, the
number and type of livestock attacked, and whether there
had been sightings of predated animals. If carcasses were
found, we asked for further details including the parts
consumed, age and sex, date and locale, what happened
after the attack, and other observations. To determine if the
livestock death was due to attacks by pumas, respondents
usually examined the carcass or injured animal for internal
and external injuries (bites and scratches), especially in the
skull, neck, and limbs. Pumas generally bite prey at the
throat, resulting in death by asphyxiation. Occasionally,
they can cover the carcass with leaves and other materials
to hide it from other predators (Hoogesteijn 2005). How-
ever, we were very confident in the respondents’ ability to
distinguish attacks by puma, we discarded unreliable
reports to eliminate any biases in our study. To evaluate
local opinion, we recorded respondents’ suggestions for
reducing losses and their feelings regarding the presence of
pumas in the area.
Data Analysis
To evaluate whether opinion on how to solve predation was
associated with factors such as respondent profile, herd
size, and history of losses, we used Pearson’s v2 analysis
(v2; P \ 0.05), which is especially suited for nominal data
(Zar 2010). To calculate the test, we arranged two-
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Environmental Management
dimensional contingency tables considering the absolute
frequencies (n) for each response (improving husbandry,
removing pumas, killing pumas, no appropriate action, and
would not answer) in relation to the age class of the
respondent, the herd size, and the history of losses
(Table 2).
To assess conflict among respondents’ opinions, we
adopted the potential for conflict index (PCI2), which is a
descriptive statistical method to facilitate the understanding
of results in human dimensions research (Manfredo et al.
2003; Vaske et al. 2006, 2010). PCI2 ranges from 0
(minimum potential for conflict) to 1 (maximum potential
for conflict). The maximum potential for conflict
(PCI2 = 1) occurs when 50 % agree, 50 % disagree, and
0 % are neutral. In contrast, no potential for conflict
(PCI2 = 0) occurs when 100 % agree, disagree, or are
neutral (Vaske et al. 2010). To calculate the PCI2, we
reorganized five dichotomous questions (yes, no, or nei-
ther) of the semistructured questionnaire (Table 2) into a
bipolar scale, with scale widths from -1 (minimum scale
value) to 1 (maximum scale value) and the distance func-
tion D2, which includes a neutral value. Power function
and number of repetitions for the simulation were accord-
ing to default values, 1 and 400, respectively. We com-
puted the frequency (n) and scale mean for each variable
among each group of respondents. Therefore, we compared
statistical differences (d) among respondents with and
without prior history of l losses, where d [ 1.96 is signif-
icant at P \ 0.05 (Vaske et al. 2006). We displayed the
results as bubble graphs to illustrate dispersion, central
tendency and form simultaneously, where large bubbles
reflect more potential conflict and small indicate less
(Manfredo et al. 2003). To calculate the PCI2, we used the
standalone online version (PCI2sa) and plotted the bubble
graph using Microsoft PowerPoint templates (http://war
nercnr.colostate.edu/^jerryv/PCI2/index.htm).
To reduce potential bias due to differences in memory of
respondents, we only quantified the frequencies (n), per-
centages (%), mean (x), and standard deviation (SD) of
livestock attacked more recently (2004–2007) as per sim-
ilar surveys (Oli et al. 1994; Palmeira and Barrella 2007;
Soto-Shoender and Giuliano 2011).
To determine when livestock predation (response vari-
able) occurred in relation to some possible explanatory
variables, we used a logistic regression model [generalized
linear model (GLM)] for a binomial distribution (Soto-
Shoender and Giuliano 2011; Teichman et al. 2013). We
created a binary history of predation (1 = presence and
0 = absence) for each respondent (n = 42) considering all
occurrences (even before 2004). As explanatory variables
(Table 2), we used the property profile (area and herd sizes
per respondent) and extracted landscape covariates such as
elevation, tree cover, and Euclidian distance from the
Environmental Management

Table 1 Semistructured questionnaire used to interview livestock breeders about puma predation on their livestock

Property and respondent profile

1. Name of property owner:

2. Property size (ha):

3. Main house (UTM): X/Y Elevation (m): GPS ID:

4. Property purposes: ( ) subsistence ( ) holiday home ( ) tourism ( ) animal ( ) crop ( ) others?

5. Name of respondent:

6. Age:

7. Place of birth:

8. Time of residence at the property:

9. Main activity:

10. Position: ( ) owner ( ) employee ( ) others?

Livestock predation

11. Number and type of livestock reared:

( ) cattle ( ) horses ( ) goats ( ) sheep ( ) pigs ( ) poultry ( ) dogs ( ) others?

12. Husbandry practices:

( ) extensive ( ) nightly confinement ( ) seasonal confinement ( ) total confinement ( ) others?

13. Did already have livestock losses due to puma? ( ) Y ( ) N

14. Number and type of livestock attacked by puma:

Date Livestock Age Gender 1234* Parts consumed Kill site (X/Y) Elevation (m)

* Livestock conditions: (1) injured (2) killed/covered (3) killed/uncovered (4) others?

15. What did you do after the attack? ( ) changed husbandry practices ( ) nothing ( ) others?

16. Did you see the puma? ( ) Y/Date/Site? ( ) N

17. Do you know someone or someplace that had livestock losses duo to puma? ( ) Y/Who/Where? ( )

Perception

18. What kind of big cats exist on the property? ( ) puma ( ) jaguar ( ) others?

19. Do you know the puma? (show photo) ( )Y( )N

20. What kind of ecological role of puma or for what puma exist? ( ) preserve/control ( ) prettify ( ) no

role ( ) don’t know ( ) others?

21. What do you think of puma close to where you live? ( ) like ( ) dislike ( ) indifferent ( ) don’t

know ( ) others?

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Table 1 continued

Perception

22. Where the most suitable place for puma to live? ( ) CJSP ( ) forest ( ) zoo ( ) don’t know ( )

others?

23. Puma poses a threat to livestock? ( ) Y/Why? ( ) N

24. Puma poses a threat to humans? ( ) Y/Why? ( ) N

25. Could you suggest how to solve livestock losses by puma predation? ( ) improve husbandry practices

( ) financial compensation ( ) remove the problem puma ( ) kill the puma ( ) don’t know ( ) others?

26. Who should bear responsibility for the damage caused by puma predation ? ( ) government ( )

livestock owner ( ) don’t know ( ) others?

27. Do you think that is necessary to control livestock predation? ( ) Y ( ) N

28. Would you change husbandry practices to reduce losses due to puma? ( ) Y ( ) N

29. Would you accept suggestions to prevent losses? ( ) Y ( ) N

30. Would like to receive the results and/or other materials from this survey? ( ) Y/Address: ( ) N

largest urban center in the study area (Municipality of
Campos do Jordão) using ArcGis 9.3 (ESRI 2008). To
avoid colinearity between the explanatory variables, we
calculated a Pearson’s correlation coefficient (r; P \ 0.05)
(Zar 2010) before adding them to the model. After this
step, we excluded the percent of tree cover from the models
because it was associated positively (r = 0.47; P \ 0.01)
with elevation. Variable elevation was selected over per-
cent tree cover because there is a considerable number of
properties located in the high-altitude grasslands where the
percent of tree cover is lower or absent and livestock
predation occurs as well. Then, the explanatory variables
were standardized to a mean of zero and unit variance. We
tested the overall model fit regarding a subset of regression

Table 2 Variables selected for

Variable description Range Value Analyses
the analyses
Tree cover (%) 27–75 Continuous GLM
Elevation (m) 1012–1985 Continuous GLM
Distance from the urban center (m) 6771–29,850 Continuous GLM
Area size (ha) 1–8341 Continuous GLM
Herd size 1–960 Continuous GLM
Herd size \20 Categorical v2
21–60
[61
Age class 18–35 Categorical v2
36–53
[54
History of livestock attacks Yes or no Categorical v2
Puma poses a threat to livestock Agree, disagree or neutral Scale (-1 to 1) PCI2
Puma poses a threat to humans Agree, disagree or neutral Scale (-1 to 1) PCI2
It is necessary to control depredation Agree, disagree or neutral Scale (-1 to 1) PCI2
Would change husbandry practices Agree, disagree or neutral Scale (-1 to 1) PCI2
Would accept suggestions to prevent losses Agree, disagree or neutral Scale (-1 to 1) PCI2
GLM generalized linear models, v2 Chi square, PCI2 potential for conflict index

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Environmental Management
parameters using the maximum likelihood estimate and
Akaike’s information criterion (AIC). We developed a
priori a set of candidate models including a global model
with all parameters (K = 5) and with the link function
‘logit.’ We used the ‘‘corrected’’ AIC (AICc) a second-
order AIC for small samples, the top delta AICc (DAICc
proximity to zero) and greater AICc weights (w) to select
the best-fit model (Burnham and Anderson 2002). We used
the inverse link function ‘plogis’ to interpret the parameters
and residuals deviance (variance and mean). We also
examined overdispersion (/) from the global model where
c-hat [1 (dispersion parameter) indicates more variance
than would be expected. Values much higher than 1 ([4)
probably indicate a lack-of-fit (Burnham and Anderson
2002; Mazerolle 2013). When variances are greater than 1,
the Quasi-likelihood (QAICc) models should adjust the
expected variance to account for / (Bolker 2008). To test
the GLM, Pearson’s correlation, and v2 analyses, we used
the package ‘stats’ in the R environment version 3.0.2
(2013-09-25; R Core Team 2013). Also, to rank the models
based on AIC, we used the package ‘AICcmodavg’
(Mazerolle 2013) and plotted the best-fit model in the
‘graphics’ (R Core Team 2013).
Results
Respondent Profile
The 36 visited properties covered a total area of
17,380.40 ha. Sixty-four percent of them ranged in size
from 1 to 84 ha and 30.5 % from 144 to 780 ha. Only
5.5 % of properties covered large areas, in particular, the
CJSP totaling 8341 ha and a sheep ranch totaling approx-
imately 4800 ha. Property purposes were subsistence
agriculture (30.5 %), followed by holiday homes (14 %),
holiday homes with crop or animal production (14 %),
tourism with animal production (14 %), animal production
(11 %), crop production (5.5 %), and other activities (the
CJSP, the Brazilian army, a monastery, and a real estate
development; 11 %).
The 42 respondents were 93 % male and 7 % female.
The mean age was 45 ± 15 years, with a minimum of 18
and maximum of 72 years. The average time of residence
on the current property was 31 ± 21 years, with a mini-
mum of one and maximum of 65 years. About 90 % of the
respondents were born in municipalities inserted in the
APA. The main activity of the respondents was classified
into general service (homemaker and housekeeper; 43 %),
animal husbandry (17 %), technical service (14 %),
administrative service (9.5 %), smallholder (7 %), and
others (retired, military, monk, and public servant; 9.5 %).
Local Opinion
When respondents were asked what kind of big cats exist
on the property, the puma was the species most frequently
cited (74 %) followed by the ocelot (Leopardus pardalis;
21 %), the jaguar (12 %), and other small felids (Leopar-
dus spp.; 5 %). When asked about the ecological role of
puma, 55 % would not answer, 19 % said preserving the
ecosystem or controlling native prey, 19 % said attacking
livestock or causing damage, and 7 % said no role. In
relation to what they thought of puma close to where they
live, 38 % dislike or feel fear, 38 % are indifferent, 19 %
like, and 5 % would not answer. When we asked where
would be the most suitable place for puma to live, 62 %
said the forest or the jungle, 14 % would not answer, 12 %
said the CJSP, 7 % said no place, and 5 % said the zoo.
When we asked who should bear responsibility for the
damage caused by predation (i.e., pay for the damages),
33 % thought the government, 31 % said the livestock
owner, 24 % said no one, and 12 % would not answer.
Thirty-three percent of the respondents would not
answer the question about how to solve livestock losses,
26 % cited that improving livestock husbandry practices
would be the best solution, 19 % said no appropriate
action, 17 % said removing the ‘‘problem’’ individual from
the area, and 5 % said killing pumas. Note that these last
three suggestions were cited after 24 % of respondents
stated ‘‘cannot kill.’’ Specific husbandry practices were
mentioned by respondents, for example, monitoring the
border, prevention, protection, fences, and confines, using
collar bells around the necks of domestic animals, intro-
ducing horned bulls into the herds, and maintaining wild
populations of preferred prey species that pumas like to eat.
The opinion on how to solve the predation problem was
independent of herd size (v2 = 2.53, df = 4, P = 0.63)
and history of losses (v2 = 1.65, df = 2, P = 0.43), and
correlated with respondent age class (v2 = 13.23, df = 4,
P = 0.01). Respondents up to 35 years tended to suggest
improving livestock husbandry practices, whereas respon-
dents older than 54 years tended to suggest removing the
‘‘problem’’ individual from the area or killing pumas. Most
of the intermediate age class between 36 and 53 years
refused to answer questions on how to solve the predation.
The PCI2 for attitudes of the respondents demonstrated
high discordance among dichotomous responses (Fig. 2).
The questions ‘‘do pumas pose a threat to livestock?’’, ‘‘do
pumas pose a threat to humans?’’ and ‘‘would you change
husbandry practices?’’ represented more potential conflict
due to the extreme values of favorable (*50 %) and
unfavorable (*50 %) responses. To the questions ‘‘is it
necessary to control predation?’’ and ‘‘would you accept
suggestions to prevent losses?’’, the responses were more
strongly in agreement (*71 %) than disagreement and
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Fig. 2 Potential for conflict

index (PCI2) for attitudes
among livestock breeders
toward livestock predation by
pumas. PCI2 ranges from 0 (no
conflict) to 1 (maximum
conflict) and is represented by
bubble size. Small bubbles
indicate less potential conflict
among respondents and large
reflect more conflict

reflected less potential conflict. There was no difference The reported losses were separated into three groups: the
among respondents with and without history of losses presence of dead livestock (60 %), disappearance of ani-
because the differences in observed PCI2 values were not mals (36 %), and injured animals (4 %). According to
statistically significant (d \ 1.96; P [ 0.05). reports, the carcasses bore marks typical of puma attacks,
such as scratches on the neck and back. The parts most
Livestock Predation commonly eaten were head, neck, chest, ribs, groin, back,
and leg. It was also very common for the predator to drink
The most numerous livestock were sheep, cattle, poultry, the blood and separate the viscera to consume the liver and
and horses, followed by pigs and goats (Table 3). The heart. Of the total livestock found dead, 17 were uncovered
distribution of the type and number of livestock by and 11 were covered by foliage. The owners only consid-
respondent was not uniform. Although many kept horses, ered disappearances as being predation events when a loss
poultry, and cattle, few respondents kept pigs, sheep, and occurred simultaneously in neighboring properties and the
goats. The average number of livestock per respondent was animal remains were visually examined.
61 ± 147, with a minimum of one and maximum of 960 Half of the respondents (50 %) reported that they had
animals. Extensive farming was used most frequently, lost some type of livestock due to pumas. Considering only
followed by semiextensive (e.g., nightly confinement), and the most recent events (2004–2007), less than half (38 %)
intensive (total confinement). When necessary, 26 % of suffered losses (Fig. 1). The average livestock loss reported
respondents used more than one type of husbandry in per respondent was 1 ± 2 (n = 42), while the average only
accordance with the type of livestock, time of year, for respondents who had losses (n = 16) was 3 ± 3, with a
financial need, and other factors. minimum of 1 and maximum of 11 animals. Following the
respondents who have livestock losses, nine owners did
nothing, four changed their herd husbandry practices, and
Table 3 Number of losses attributed to pumas from 2004 to 2007
and herd size estimated in 2007 three buried the carcasses of the animals killed.
Predation on horses was most commonly reported
Livestock 2004 2005 2006 2007 Total Herd size (2007)
(51 %), followed by cattle (28 %), sheep (17 %), and goats
Horses 5 4 10 5 24 279 (4 %), totaling 47 animals attacked (Table 3). Mean annual
Cattle 1 1 3 8 13 695 predation was approximately 12 ± 5 animals attacked,
Sheep 1 2 3 2 8 860 equivalent to 0.4 % of the total livestock at the properties.
Goats 1 0 1 0 2 4 Of the animals attacked, 57 % were males, 30 % females,
Pigs 0 0 0 0 0 36 and the gender of 13 % could not be determined, often
Poultry 0 0 0 0 0 689 because they were newborn animals without sexual iden-
Total 8 7 17 15 47 2563 tification. Regarding age group, only 32 % animals had
their age estimated, eight of which were up to 1 year, four

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Table 4 Generalized linear

Candidate Models K AICc DAICc w
models (GLM) for determining
the probability of livestock Landscape (elevation ? distance from the urban center) 3 56.51 0.00 0.44
predation on the site covariates
Landscape (elevation ? distance from the urban center) ? herd size 4 58.14 1.63 0.19
Landscape (elevation ? distance from the urban center) ? area size 4 58.56 2.05 0.16
Constant model (*parameters) 1 60.32 3.81 0.06
Global model (all parameters) 5 60.41 3.89 0.06
Property profile (area ? herd sizes) ? elevation 4 61.14 4.63 0.04
Property profile (area ? herd sizes) 3 62.10 5.59 0.03
Property profile (area ? herd sizes) ? distance from the urban center 4 62.90 6.39 0.02
K number of parameters, AICc ‘‘corrected’’ Akaike’s information criterion for small samples, DAICc delta
AICc differences, w Akaike weights

elevations was demonstrated using the best-fit model

(Fig. 3).

Discussion

In our interviews, there was a high non-response rate,

Fig. 3 Probability of livestock predation in relation to elevation and
distance from the urban center of the best-fit generalized linear model
(GLM)
2 years, and three over 10 years old. For almost half of the
losses (45 %), the exact time when the attacks occurred
could not be identified. The remainder occurred during the
rainy season between spring and summer (September–
January; 34 %) and the dry season that precedes the winter
(April, May, and June; 21 %).
Based on the top DAICc (proximity to zero) and w, the
supported model indicated that predation was expected to
be higher in areas far from the urban center and at high
elevations (Table 4). There was a moderate / in the global
model because the variance was slightly greater than 1 (c-
hat = 1.31), and for this reason, we opted for AICc rather
than QAICc. The selected model had K = 3,
DAICc = 0.00, w = 0.44. The model also had good pre-
dictive power (mean P = 0.46, range 0.16–0.84), and
explained 49.88 % of the deviance. Area and herd sizes
were weak predictors of the probability of livestock pre-
dation in the models. The probability of predation occur-
ring in areas far from the urban center and at high
especially for the question on how to solve livestock losses.
As demonstrated in the results, many respondents stated
‘‘cannot kill’’ before answering the question, suggesting
that they were fearful of being reported to the authorities if
they suggested that problem pumas should be killed, even
after we promised anonymity and confidentiality on the
questionnaire application. The principal implication of the
non-responses was the lack of consensus among livestock
breeders and an increase in the potential for conflict. With
the exception of respondent age, the opinions on how to
solve predation by pumas were independent of the
respondent profile, the characteristics of the property, or
the occurrence of losses. In general, opinions regarding
livestock predation by pumas and jaguars do not seem to be
associated with the respondent profile, herd size, number of
losses (Palmeira and Barrella 2007), or size of the property
(Conforti and Azevedo 2003; Zimmermann et al. 2005).
Occasionally, the opinion is associated with the number of
livestock held by respondents (Conforti and Azevedo
2003), the history of attacks (Soto-Shoender and Main
2013), and the age of respondents (Zimmermann et al.
2005).
The majority of attitudes toward predation suggested
high potential for conflict among the respondents in the
survey. The questions ‘‘do pumas pose a threat to livestock
or humans?’’ and ‘‘would you change husbandry prac-
tices?’’ represented higher potential for conflict because of
the lack of consensus among livestock breeders. Puma
attacks on humans are rare in Brazil, and despite there
being no record of fatal attacks in the study area, half of
livestock breeders reported that the species poses a threat to
humans as well as to their livestock. In contrast, livestock

123

breeders tended to be in agreement with the questions ‘‘is it
necessary to control predation?’’, representing a moderate
conflict level, and ‘‘would you accept suggestions to pre-
vent losses?’’ being the lowest conflict level. There was a
discord between high consensus to accept suggestions to
prevent losses and little consensus to change husbandry
practices. The results of the potential for conflict should be
used to guide implementation of mitigation measures and
demonstrate a critical need to obtain a consensus of opinion
before implementing management actions.
Predation was low and represented only 0.4 % of the
livestock held in the study area. Other surveys have also
shown that the number of losses attributable to predator
attacks is relatively low and generally does not exceed 3 %
of the total stock (Graham et al. 2005). We believe that
although pumas might not cause all livestock disappear-
ances, these reported disappearances play an important role
in human–wildlife interactions as respondents consider
predators responsible. Commonly, livestock disappear-
ances with no carcasses being found have also represented
a large proportion of losses reported by herders and tend to
be blamed on predators (Palmeira and Barrella 2007;
Rosas-Rosas et al. 2008).
Horses were the most frequently attacked livestock,
and although well distributed spatially, they did not rep-
resent the most abundant livestock in the study area,
which indicates that predation was selective and influ-
enced by the preference of the predator. This fact has
been well reported among wild carnivores that attack
certain types of livestock more than expected (Namgail
et al. 2007; Palmeira and Barrella 2007; Azevedo 2008;
Zarco-González et al. 2012). Pumas attacking horses are
seldom reported in the literature; they usually attack foals
and young horses but rarely adult horses (Graipel et al.
2004; Shaw et al. 2007; Gray et al. 2008). When large
prey like buffalo (Azevedo et al. 2010) or cattle are
attacked, these are generally young or newborn individ-
uals (Polisar et al. 2003; Palmeira et al. 2008). This trend
was also found in our study as the horses and cattle that
were attacked were mostly juveniles. Permanent confine-
ment of juveniles and nightly confinement of adults
should be implemented as alternative husbandry practices
to mitigate this predation.
The livestock losses with a confirmed date occurred
during two distinct periods: in the rainy season between
spring and summer and in the dry season that precedes
winter. The occurrence of livestock predation by carnivores
in specific periods seems to be very common and has been
observed both in winter (Palmeira and Barrella 2007) and
in summer, and during the wet season or dry season,
depending on the region (Patterson et al. 2004; Kolowski
and Holekamp 2006; Palmeira et al. 2008; Li et al. 2013).
These peaks of predation in certain seasons may be
123
Environmental Management
associated with seasonal birth of livestock (Palmeira et al.
2008) or the availability of alternative prey at different
times of the year (Polisar et al. 2003; Patterson et al. 2004;
Kolowski and Holekamp 2006).
Although extensive management of livestock was the
most frequently used management method, respondents
employed more than one type of husbandry simultaneously
depending on need. Synchronized husbandry seems to be a
very common practice among livestock breeders (Palmeira
and Barrella 2007), which limits our ability to investigate
further the relationship between the livestock husbandry
practices adopted and the number of losses. Nevertheless,
some studies have suggested that predation is strongly
associated with husbandry practices adopted by farmers,
and that changes could reduce some of the attacks (Polisar
et al. 2003; Ogada et al. 2003; Wilson et al. 2006; Li et al.
2013). A simulation using three different management
scenarios concluded that adopting a rotation of seasonal
breeding could reduce the exposure of animals to pumas
(Kissling et al. 2009). In other studies, the losses were not
associated with the type of husbandry adopted (Graham
et al. 2005), the size or structure of the pens, or the number
of guard dogs or herders (Hemson et al. 2009).
Elevation and distance from the urban center were
important predictors of the supported model. Areas far
from the urban center and at high elevations are more
likely to suffer losses to pumas because of the habitat
suitability for the species. Part of this population is found
only in original habitat remnants above 800 m (Mazzolli
1993). In British Columbia, cougar–human conflicts were
more likely to occur close to roads and at intermediate
elevations (Teichman et al. 2013). In Mexico, predation
was correlated with vegetation type and distances to per-
manent water and roads (Rosas-Rosas et al. 2010). In
Guatemala, forest cover, distance to forest, bodies of water
and human settlements were also landscape predictors of
the probability of livestock predation (Soto-Shoender and
Giuliano 2011).
Despite the low rates of predation, some preventive
strategies must be addressed to reduce livestock losses.
Managing problem animals such as killing or re-locating
offending pumas might not be possible because the
Atlantic forest is an endangered biome where there is
evidence of a genetic bottleneck for pumas (Miotto et al.
2011; Castilho et al. 2012). Even if the problem animal is
detected, the efforts to capture are enormous and translo-
cation appears to have a low success rate (Linnell et al.
1997). In the same way, removing problem individuals will
not work in multi-use landscapes where livestock are dis-
persed throughout the same areas in which carnivores are
to be conserved (Linnell et al. 1999). Also, financial
compensation for those suffering losses is not a solution to
the problem of conserving large carnivores (Dickman et al.
Environmental Management
2011) and can in fact decrease the efforts of livestock
breeders to prevent losses and exacerbate conflicts with
wildlife (Bulte and Rondeau 2005). Therefore, more
financial incentives are necessary for husbandry practices
that reduce probability of attacks, such as nightly con-
finement for horses, sheep and goats, and keeping calves
away from the sites inhabited by big cats. Financial
incentives could also be applied for intensive husbandry
practices that improve livestock production as well as
finding sustainable, wildlife friendly and compatible
alternatives to supplementing their income.
Developing environmental education and offering
technical support for husbandry practices should be further
developed, because most respondents reported that they
would be able to accept suggestions to prevent livestock
losses, which represents the lowest potential for conflict.
Actions of environmental education should be compre-
hensive and provide knowledge regarding a broad range of
topics on biology, ecology, and conservation (e.g., eco-
logical role of predators and natural prey, their conserva-
tion significance, and intrinsic value). These actions could
be performed by a trained team of professionals in social
and biological sciences and directed to children, key-in-
formants, and information disseminators (e.g., teachers,
monitors and park guards, tourist guides, and volunteers).
The activities should occur in the field or classroom using
some concepts of the constructivism theory where humans
can be active learners through their experiences. A specific
environmental education program must be directed to
livestock breeders incorporating technical information on
alternative husbandry practices to improve animal pro-
duction and, consequently, reduce the risk of livestock
losses and the possible retaliatory hunting of predators.
Also, it should be necessary to monitor losses to the
properties inserted in a landscape located in areas with high
probability of predation, particularly those properties situ-
ated far from urban centers and at high elevations.
This study’s recommendations were devised by con-
servation biologists in collaboration with an agronomist
specialized in animal production and should be directed to
a broad range of stakeholders such as livestock breeders,
wildlife conservationists, and political or decision-makers.
It is important to emphasize that our inference to a broader
and more general context and scope comes from a small
sample size, and the results and conclusions only apply to
the study area. Because predictors of livestock predation
should vary among sites, the choice of management actions
would also depend on the region. This survey represented
the first attempt in the study of livestock predation to
evaluate potential for conflict among livestock breeders
regarding predators and it is the main contribution and
novelty of this study.
Acknowledgments Thanks to the Horto Florestal (Campos do
Jordão State Park), especially Marco Antonio P. Marcondes and
Waldir Joel de Andrade, and the Instituto Florestal (Process SMA N
42.769/2006) for institutional support and permission to conduct this
survey. Thanks to all the livestock breeders who kindly agreed to
participate in the interviews, especially Seu Jorge, Pedro Henrique de
Oliveira, Fabinho, and Nelson M. Marcelino for their field assistance
in data collection. We thank the Environmental/Forest Police and Fire
Department at Campos do Jordão for information provided, as well as
Pedro Paulo Filho, Arthur Bicudo, Antonio Scafuto Neto, Orlando
Paggiaro, Mr. Roberto (Romalar), Fotótica Shimazu, Fernando, Tião
and Cali for donation of photos and other materials. We thank our
parents, Ana Maria T. Trinca, Walter Trinca, and Benvinda B. Lopes
for the infrastructure provided and complete financial support of
fieldwork activities. Thanks to Ronaldo G. Morato for a reference
provided and Renata A. Miotto for helping us to find the target
journal. We are very grateful for helpful comments and suggestions
from reviewers, Heather Aslin and José R. Soto-Shoender, and the
editors-in-chief.
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